Mediterranean Marine Science

Vol. 22, 2021

Species and habitats of conservation interest in the Ecologically and Biologically Significant Area of the Strait of Sicily: a contribution towards the creation of a Specially Protected Areas of Mediterranean Importance

CONSOLI PIERPAOLO Zoological Station Anton Dohrn (SZN), Department of Integrative Marine Ecology, Messina, Italy & Italian Institute for Environmental Protection and Research (ISPRA), Milazzo (ME), Italy ALTOBELLI CHIARA Italian Institute for Environmental Protection and Research (ISPRA), Palermo, Italy & National Institute of Oceanography and Experimental Geophysics (OGS), Trieste, Italy PERZIA PATRIZIA Italian Institute for Environmental Protection and Research (ISPRA), Palermo, Italy BO MARZIA Università degli Studi di Genova, Dipartimento di Scienze della Terra, dell’Ambiente e della Vita, Genova, Italy ROSSO ANTONIETTA Università degli Studi di Catania, Dipartimento di Scienze Biologiche, Geologiche e Ambientali, Sezione di Scienze della Terra, Catania, Italy ALONGI GIUSEPPINA Università degli Studi di Catania, Dipartimento di Scienze Biologiche, Geologiche e Ambientali, Sezione di Biologia vegetale, Catania, Italy SERIO DONATELLA Università degli Studi di Catania, Dipartimento di Scienze Biologiche, Geologiche e Ambientali, Sezione di Biologia vegetale,

http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 06/10/2021 09:44:34 | Catania, Italy CANESE SIMONEPIETRO Zoological Station Anton Dohrn (SZN), Department of Integrative Marine Ecology, Napoli, Italy ROMEO TERESA Zoological Station Anton Dohrn (SZN), Department of Integrative Marine Ecology, Messina, Italy & Italian Institute for Environmental Protection and Research (ISPRA), Milazzo (ME), Italy ANDALORO FRANCO Italian Institute for Environmental Protection and Research (ISPRA), Palermo, Italy & Zoological Station Anton Dohrn (SZN), Department of Integrative Marine Ecology, Palermo, Italy https://doi.org/10.12681/mms.25125

To cite this article:

CONSOLI, P., ALTOBELLI, C., PERZIA, P., BO, M., ROSSO, A., ALONGI, G., SERIO, D., CANESE, S., ROMEO, T., & ANDALORO, F. (2021). Species and habitats of conservation interest in the Ecologically and Biologically Significant Area of the Strait of Sicily: a contribution towards the creation of a Specially Protected Areas of Mediterranean Importance. Mediterranean Marine Science, 22(2), 297-316. doi:https://doi.org/10.12681/mms.25125

http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 06/10/2021 09:44:35 | Research Article Mediterranean Marine Science Indexed in WoS (Web of Science, ISI Thomson) and SCOPUS The journal is available on line at http://www.medit-mar-sc.net DOI: http://dx.doi.org/10.12681/mms.25125

Pierpaolo CONSOLI1,2, Chiara ALTOBELLI3,4, Patrizia PERZIA3, Marzia BO5, Antonietta ROSSO6, Giuseppina ALONGI7, Donatella SERIO7, Simonepietro CANESE8, Teresa ROMEO1,2 and Franco ANDALORO3,9

1 Zoological Station Anton Dohrn (SZN), Department of Integrative Marine Ecology, Messina, Italy 2 Italian Institute for Environmental Protection and Research (ISPRA), Milazzo, Italy 3 Italian Institute for Environmental Protection and Research (ISPRA), Palermo, Italy 4 National Institute of Oceanography and Experimental Geophysics (OGS), Trieste, Italy 5 University of Genoa, Department of Earth, Environmental and Life Sciences, Genoa, Italy 6 University of Catania, Department of Biological, Geological and Environmental Sciences, Section of Earth Sciences, Catania, Italy 7 University of Catania, Department of Biological, Geological and Environmental Sciences, Section of Plant Biology, Catania, Italy 8 Zoological Station Anton Dohrn (SZN), Department of Integrative Marine Ecology, Napoli, Italy 9 Zoological Station Anton Dohrn (SZN), Department of Integrative Marine Ecology, Palermo, Italy

Corresponding author: [email protected]

Contributing Editor: Vasilis GEROVASILEIOU

Received: 22 Octber 2020; Accepted: 5 March 2021; Published online: 10 May 2021

Abstract

In 2015, the Strait of Sicily, which includes several banks, was candidate as a future Specially Protected Area of Mediterranean Importance (SPAMI) by the Contracting Parties of the Barcelona Convention. In this context, the present study aims to provide the first biological and ecological characterisation of this poorly known area, focusing on habitats and species of conservation concern. Surveys were carried through a remotely operated vehicle (ROV) and allowed the identification of 19 habitat types, mostly considered of priority interest by the SPA/BD Protocol of the Barcelona Convention. A total of 269 taxa were also identified, mostly Porifera, Cnidaria and Pisces, among which 115 species are of conservation concern according to the IUCN (IUCN) Red Lists, the Habitats Directive, the Bern Convention, the SPA/BD Protocol and CITES. ROV surveys allowed some interesting ob- servations on poorly known species and habitats. Results underlined that remarkable discrepancies in species conservation status assessment exist between the several lists of protected species considered. The IUCN Red Lists, although not legally binding for European States, are the most complete, but, in spite of this, the conservation policies in the EU are largely focused on the Habitats Directive, which is really not exhaustive. An exhaustive and legally binding instrument to protect species of conservation concern at European scale is highly recommended. Acquired results could be useful for the delimitation of a future SPAMI or a network of Marine Protected Areas (including the investigated banks) and the identification of zones within them suitable for different area-based management measures.

Keywords: Biodiversity; environmental protection; epibenthos; habitat; Habitats Directive; IUCN Red List; Mediterranean Sea banks; SPA/BD Protocol.

Introduction perturbations (Worm et al., 2006; Danovaro et al., 2008; Cardinale et al., 2012; Hooper et al., 2012). The Mediterranean Sea is considered a hot spot of In the last decades, the deficient conservation status of biodiversity with approximately 17,000 marine species many biota has led to initiatives aimed at halting the loss (Coll et al., 2010; Costello et al., 2010). Since the begin- of biodiversity (Ban et al., 2014; Boyes & Elliot, 2014). ning of the Anthropocene, following the industrial rev- The main international actions for the protection of coast- olution, biodiversity loss accelerated massively (Rock- al and marine habitats are the Washington Convention on ström et al., 2009a, 2009b) impacting marine ecosystem International Trade in Endangered Species of Wild Fauna services, and increasingly impairing the ocean’s capacity and Flora (CITES; UNEP, 1973), the Bern Convention to provide food, maintain water quality, and recover from (Conservation of European Wildlife and Natural Habi-

Medit. Mar. Sci., 22/2 2021, 297-316 297

http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 06/10/2021 09:44:35 | tats; Council of Europe, 1979), the United Nations Con- Vecchia Banks in the Strait of Sicily and indicated that vention on Biological Diversity (CBD; UNEP, 1992), the they represent unknown hot spots of biodiversity threat- Protocol concerning Specially Protected Areas and Bio- ened by human activities in the wide and diversified ‘Sic- logical Diversity in the Mediterranean (SPA/BD Proto- ily Channel’ EBSA. According to Altobelli et al. (2017) col) of the Barcelona Convention (UNEP/MAP, 1995) and Consoli et al. (2018), fishing, marine debris, inva- and the International Union for Conservation of Nature sive alien species and oil and gas extraction represent (IUCN, 2015). The Habitats Directive (EU, 1992) and the major pressures and threats to these banks. Using a the Birds Directive (EU, 2009), allowing the establish- semi-quantitative methodology, Altobelli et al. (2017) ment of the largest continent-wide Natura 2000 network also showed that these banks could represent eligible of Special Areas of Conservation (SACs; Hochkirch et sites for the implementation of efficient and realistic al., 2013), and the Marine Strategy Framework Direc- area-based management measures (SPAMI, MPAs), as tive (EU, 2008) are among the most important European expected by the 2015 RAC/SPA meeting to be able to Union directives on this topic. guarantee protection and sustainable use. In the Mediterranean Sea, the SPA/BD Protocol of In light of this, the main aim of the present research the Barcelona Convention (UNEP/MAP, 1995) led to the was to provide a useful baseline for the establishment of establishment of 32 Specially Protected Areas of Medi- a future SPAMI associated with these banks by assessing terranean Importance (SPAMIs), in order to promote co- species and habitat diversity with a particular focus on operation in the management and conservation of natural those of conservation concern according to the main in- areas, as well as in the protection of threatened species ternational conservation policies. and their habitats. In 2015, during the second meeting of the RAC/SPA (Regional Activity Centre for Specially Protected Areas), Material and Methods established by the Contracting Parties to the Barcelona Convention and its Protocols in order to assist Mediterra- Study area nean countries in implementing the SPA/BD Protocol, experts started a review of the existing literature regarding The term ‘banks’ will be hereafter used to indicate the Strait of Sicily (central Mediterranean Sea) in order to submarine reliefs such as shallows, ridges, knolls and assess the possibility of creating one or more SPAMIs in pinnacles. Within banks we also include seamounts, i.e., this area (RAC/SPA, 2008). The Strait of Sicily (or Sicily submarine elevations of volcanic origin occurring in the Channel), which includes several banks (submarine re- Strait of Sicily; according to the definition by Staudigel liefs such as shallows, ridges, knolls, pinnacles and sea- et al. (2010) these refer to ‘any geographically isolated mounts) of different sizes and origins, has been already topographic feature on the sea floor taller than 100 m, recognised as an Ecologically or Biologically Significant including ones whose summit regions may temporarily Area (EBSA) by the contracting parties of the Conven- emerge above sea level, but not including features that tion on Biological Diversity, due to its high naturalistic are located on continental shelves or that are part of other importance (CBD-COP 12 Decisions, 2014). major landmasses’. In general, banks are ecologically important, especial- Owing to its location, the Strait of Sicily represents ly those located close to, or falling within, the photic zone a crucial connection between the Western and Eastern because they present high productivity, biomass and bio- Mediterranean basins. Located between Italy, Tunisia diversity, usually benefitting from enhanced organic in- and Malta, it is a vast physiographic structure of about puts under low pollution conditions compared to coastal 250000 km2 oriented NW-SE between the African conti- areas. Moreover, banks are important aggregating zones nent and Sicily. It consists of continental shelves dissect- (feeding, nursery, spawning and/or recruitment areas) for ed by regional foult systems to form grabens (the Pan- many migratory fish, cetaceans, seabirds and cephalo- telleria, Malta and Linosa grabens, over 1300, 1500 and pods (Koslow, 1996; Pitcher et al., 2007; Fiorentino et 1700 m deep, respectively) and horsts elevating locally al., 2011; Kvile et al., 2014). In spite of this, banks are into numerous banks. Three of these banks, namely the among the least investigated marine areas in the world, Graham, Nereo and Pantelleria Vecchia Banks were in- due to survey difficulties related to their rough topogra- vestigated (Fig. 1). They are located in international wa- phy, their location (usually offshore), and the vigorous ters (UNCLOS, 1982) between Pantelleria Island and the current regime to which they are exposed (Bo et al., south-western coast of Sicily. 2012a). In the last few decades, however, the employ- The Graham Bank, along with Terrible Bank and Ner- ment of increasingly sophisticated remotely operated ve- ita Bank, is part of a large underwater horseshoe-shaped hicles (ROVs) has allowed more intensified exploration. volcanic relief named the Empedocle Seamount, located Non-destructive ROV surveys have been revealing the between 250 and 500 m in depth. The Graham Bank, in high complexity and ecological value of banks, as well as turn, is composed of numerous volcanic cones, mainly the human disturbances that threaten their environmen- aligned NW-SE, including the Ferdinandea Bank. These tal status (Bo et al., 2012a; 2014; Angiolillo et al., 2015; cones vary widely in both size and depth, most of them Consoli et al., 2016, 2018). having tops located at 45-80 m, except for the Ferdinan- Altobelli et al. (2017) evaluated the ecological and dea Bank, which rises to 9 m depth (Falzone et al., 2009; biological value of the Graham, Nereo and Pantelleria Würtz & Rovere, 2015).

298 Medit. Mar. Sci., 22/2 2021, 297-316

http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 06/10/2021 09:44:35 | Fig. 1: Investigated banks (square points) of the Strait of Sicily following the official delimitation by the Italian Navy Hydro- graphic Institute.

The Nereo Bank is one of the numerous shallows on a 40% lateral overlap. Multibeam data were processed by the Adventure Plateau, rising from the wide continental Caris HIPS 8.1 software. Then, species and habitat diver- shelf in the north-western sector of the Strait of Sicily sity were qualitatively investigated by video recordings (Colantoni et al., 1985). These morphological highs, in- using an ROV along 14 transects. cluding the Pantelleria Vecchia Bank, compose the Ad- The ROV (‘Pollux III’ Global Electric Italiana) was venture Archipelago. The Nereo Bank shows a very ir- equipped with several instruments, as already described regular shape and a rough morphology and consists of by Consoli et al. (2018). three main NE-trending minor shoals parallel to each In order to verify the presence of species of conserva- other and separated by narrow channels. In this study, the tion concern undetectable from images, four small sam- main northern ridge was investigated. It is about 3.7 km ples of coarse biogenic substrate were collected by the long and 3.5 km wide, ranging from about 30 m to 60 m ROV grabber in particular habitats (maërl and rhodolith in depth (Civile et al., 2008). beds) selected by visual census. The Pantelleria Vecchia Bank is composed of two main shoals, which elevate from 46-60 m to 16-24 m depth, and a number of smaller isolated bathymetric Video and data analysis highs (Lodolo & Ben-Avraham, 2015). Overall, 14 transects ranging from 46 to 1760 m length were performed at depths of 20-220 m (perpendic- Field sampling methods ular to the respective isobaths, from bank tops to greater depths) for a total distance of approximately 14 km. The Surveys were carried out in June-July 2014 and June characteristics of each transect are reported in Table 1. 2015 by the Italian Institute for Environmental Protection A digital archive providing over 20 h of high-defini- and Research (ISPRA) on board of the R/V Astrea. Sur- tion videos was created and 1630 images were obtained veys were carried out as part of the research project ‘Ob- every 10 seconds through the extraction of frames from servatory of the biodiversity of the marine and terrestrial the video tracks by means of free internet software environment of the Sicily Region’ aimed at developing (DVDVideoSoft). These images corresponded to almost monitoring techniques to assess protected species and 16500 m2 of analysed surface (each frame measuring habitats associated with the Nereo, Graham and Pantelle- about 3 m2, as estimated by ImageJ software). ria Vecchia Banks in the Strait of Sicily. Video frames and images were examined by experts First of all, the study area was mapped using a Kongs- for the identification of organisms, at the lowest taxo- berg EM2040 Multibeam echosounder (Kongsberg, Nor- nomic level possible, and habitats. way); data acquisition frequency was 300-400 kHz with As regards samples collected by ROV grabber, select-

Medit. Mar. Sci., 22/2 2021, 297-316 299

http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 06/10/2021 09:44:35 | Table 1. Summary of the 14 transects performed by the ROV. Surveyed areas per transect were calculated using the ROV track lengths (estimated by the GIS software ESRI ArcMAP 10.1) and an estimated width visibility (visual field) of 2 m. Habitats Directive (92/43/EC; EU, 1992), the Interpretation Manual of European Union Habitats – Area ID dive Year Transect Visual swept area Depth range Mean depth EUR28length (European (m) Commission,(m2) 2013),(m) and the identification(m) sheets of priority habitats according to

Graham 1 2014the SPA/BD 1387 Protocol (Relini2774 & Giaccone,80-180 2009). 113 Graham 2 2014 1123 2246 40-100 92 Lists of species from the studied banks (observed through the ROV video recordings and collected Graham 3 2014 916 1832 120-220 153 Graham 4 2014through 987 ROV grabber)1974 were compared100-180 with those reported 75 in the Habitats Directive, and several Graham 5 2014 46 92 80-90 82 Graham 6 2014other international 1250 conventions2500 and protocols70-170 (i.e., CITES, 96 Bern Convention, SPA/BD Protocol of Graham 7 2015 484 968 20-120 65 the Barcelona Convention), in order to identify protected or endangered species or those whose Graham 8 2015 1310 2620 80-200 150 Nereo 9 2014exploitation 925 is regulated.1850 35-45 40 Nereo 10 2014 882 1764 25-50 38 Nereo 11 2014Furthermore, 1600 in order 3200to include a larger30-60 number of 47species of conservation concern, the status of Pantelleria Vecchia 12 2014each species 879 and its relative1758 Population20-51 Trend (Decreasing34 Population Trend – DPT, or Unknown Pantelleria Vecchia 13 2014 1760 3520 20-52 35 Pantelleria Vecchia 14 2015Population 713 Trend – UPT)1426 was assessed20-60 using the IUCN43 Red List (not legally binding for European Total 14262 28524 member states) at Global, Mediterranean (IUCN, 2015) and National level (Italian Committee of ed dry and uncleaned specimens of Bryozoa were exam- Trend – DPT, or Unknown Population Trend – UPT) was ined under a scanning electron microscope (SEM)IUCN and,, 2016). assessed using the IUCN Red List (not legally binding for Corallinales, species were identified according to the for European member states) at Global, Mediterranean standard methodology (see Alongi et al., 2002).The Taxon IUCN- (IUCN,Red Lists 2015) were and also National used level to calculate (Italian Committee the Red Listof Index (RLI; see Moser et al., 2016) omy and nomenclature, as well as the status assessment IUCN, 2016). for bryozoans, followed Rosso & Di Martino (2016)for the and taxonomic The IUCNgroups Red of ListsCnidaria were andalso Pisces.used to calculateThe RLI the measures the rate of biodiversity loss; it references therein. Red List Index (RLI; see Moser et al., 2016) for the taxonomic groups of Cnidaria and Pisces. The RLI measures is an index theof therate proportionof biodiversity of loss;species it is expectedan index of to the remain propor -extant in the near future in the absence List of habitats, species and their conservation status tion of species expected to remain extant in the near fu- of any conservationture in the action absence (Butchart, of any conservation 2007). action (Butchart, The habitat types, including biocoenoses, associations 2007). and facies (Pérès & Picard, 1964) and their conservationThe revised RLIThe is revised defined RLI as: is defined as: status, were assessed following the Annex I – ‘Natural habitat types of Community Interest whose conservation ( , ) requires the designation of Special Areas of Conserva- 1 (1) RLIt = 𝑁𝑁𝑁𝑁 tion’ of the EU Habitats Directive (92/43/EC; EU, 1992), ∑𝑠𝑠𝑠𝑠=1 𝑊𝑊𝑊𝑊 𝑐𝑐𝑐𝑐 𝑡𝑡𝑡𝑡 𝑠𝑠𝑠𝑠 the Interpretation Manual of European Union Habitats – − 𝑊𝑊𝑊𝑊𝐸𝐸𝐸𝐸𝐸𝐸𝐸𝐸 ∗ 𝑁𝑁𝑁𝑁 EUR28 (European Commission, 2013), and the identifi- where ‘t’ is the year; ‘Σs’ is a summation over all assessed cation sheets of priority habitats according to the SPA/ non-Data Deficient species; ‘Wc(t,s)’ is the weight of BD Protocol (Relini & Giaccone, 2009). category ‘c’ assigned to species in year ‘t’ (weight rang-

Lists of species from the studied banks where(observed ‘t’ ises the from year; 0 – ‘ ΣLeasts’ is Concern a summation to 5 – Extinct);over all ‘WassessedEX’ is the non -Data Deficient species; ‘Wc(t,s)’ is through the ROV video recordings and collected through weight assigned to Extinct species (given the value 5 us- ROV grabber) were compared with those reportedthe weightin the ofing category the recommended ‘c’ assigned ‘equal to steps’species weights, in year with ‘t’ Criti(weight- ranges from 0 – Least Concern to 5 Habitats Directive, and several other international con- cally Endangered = 4, Endangered = 3, Vulnerable = 2, ventions and protocols (i.e., CITES, Bern Convention, Near Threatened = 1, Least Concern = 0); and ‘N’ is the SPA/BD Protocol of the Barcelona Convention),– inExtinct); order total‘W numberEX’ is ofthe assessed weight species, assigned excluding to thoseExtinct consid species- (given the value 5 using the to identify protected or endangered species or those spe- ered Data Deficient and those assessed as Extinct in the cies whose exploitation is regulated. recommendedyear the‘equal set of speciessteps’ was weights, first assessed. with For speciesCritically de- Endangered = 4, Endangered = 3, Furthermore, in order to include a larger number of fined as ‘Data Deficient’ or ‘Not Evaluated’ in the Italian species of conservation concern, the status of eachVulnerable species Red= 2, List, Near we Threatened looked at the = category 1, Least indicated Concern in the = 0);Med and- ‘N’ is the total number of assessed and its relative Population Trend (Decreasing Population iterranean list; if the species was still ‘Not Evaluated’, we 8 300 Medit. Mar. Sci., 22/2 2021, 297-316

http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 06/10/2021 09:44:35 | searched the Global list. The species was then considered was possible to distinguish 97 taxa of Porifera, only 42% ‘Not Evaluated’ or ‘Data Deficient’ when this status was were identified at species level through visual census. present in all three lists. For Cnidaria, 63 taxa were distinguished, 51 of which at RLI values can vary from 0 (all species are Extinct) to species level (81%) and for Pisces, 51 species out of 54 1 (all species are of Least Concern). taxa (95%). It was also possible to identify 13 species of Echinodermata and 10 species of Macrophyta. Moreover, the analysis of samples collected through Results ROV grabber in “maërl and rhodolith beds” led to the identification of 13 taxa of calcareous Rhodophyceae Census of habitat types forming free-living biogenic concretions, and 68 taxa of associated Bryozoa (Appendix S1). The ROV survey along the transects on the Graham, The visual census allowed identification of 115 taxa Nereo and Pantelleria Vecchia Banks (Figs 2-4) allowed of conservation concern, i.e., whose conservation or ex- identification of a total of 19 habitat types (17 if ‘associ- ploitation is regulated by several international directives ations with rhodoliths and maërl’ are counted as a single and conventions (Table 3): 108 taxa, mostly represented habitat type), including biocoenoses, associations and by Cnidaria and Pisces, are listed in the Global, Mediter- facies (Table 2). According to the Interpretation Man- ranean (IUCN, 2015) and Italian (Italian Committee of ual of European Union Habitats – EUR28 (European IUCN, 2016) IUCN Red Lists, 22 in the SPA/BD pro- Commission, 2013), 13 of them belong to three marine tocols, 16 in the Bern Convention, 10 in the CITES and habitat types (‘1120-Posidonia oceanica meadows’, only four are included in the Habitats Directive. ‘1170-Reefs’, and ‘8330-Submerged or partially sub- Twenty-two taxa out of the 108 listed in the IUCN merged sea caves’) included in the Annex I of the EU Red List fell into threatened categories, although to dif- Habitats Directive. In particular, ‘Posidonia meadows’ ferent degrees: 1 ‘Critically Endangered’ (Funiculina are considered a priority natural habitat type by the Habi- quadrangularis), according to the Italian IUCN Red List, tats Directive. Moreover, according to the SPA/BD Proto- 7 ‘Endangered’, and 17 ‘Vulnerable’ (Fig. 7). Only Cor- col (Barcelona Convention), 14 of the 19 registered hab- allium rubrum, which is considered as Endangered in the itats are considered of priority interest (habitats whose Mediterranean and Italian Red Lists, is included in the conservation is mandatory), whereas four are listed as Habitats Directive, Annex V regarding species of ‘Com- remarkable (habitats that deserve specific attention or munity interest whose taking in the wild and exploitation management). The SPA/BD classification system takes may be subject to management measures’. Most of the 22 into account five habitat variables: vulnerability, rarity, taxa falling into threatened categories (V, E, CE) showed heritage value, aesthetic value and economic value, each a decreasing population trend. The RLI was calculated of which is evaluated on three levels (high, medium and for Pisces and Cnidaria, the taxonomic groups with the low). On the basis of the above-mentioned habitat vari- highest number of taxa identified at species level listed ables, habitats are classified into three main categories in the IUCN Red Lists. For Pisces the RLI was 0.97 and (P: Priority; R: Remarkable; OH: Other Habitat) (Relini for Cnidaria 0.89. & Giaccone, 2009; Fig. 5). In the study area, the ‘associ- The analysis of samples collected by ROV grabber led ation with rhodoliths’ (IV.2.2.1.) within the ‘biocoenosis to the identification of two calcareous Rhodophyceae of of the coastal detritic bottom’ is the only habitat listed as conservation concern, namely Phymatolithon calcareum OH. It is important to note that according to the SPA/BD and Lithothamnion corallioides, included in the Annex Protocol the ‘association with rhodoliths’ and the ‘facies IV of the Habitats Directive (Table 3). with maërl’ are found in two distinct biocoenoses (i.e., The endemic ecological status is a relevant parameter ‘biocoenosis of coarse sands and fine gravels under the to assess whether a species is of ‘Community Interest’ influence of bottom currents’, and ‘biocoenosis of coastal according to the Habitats Directive and then listed in the detritic bottom’), and their classifications change from P Annex II and/or Annex IV or V. Six endemic species of to OH and from P to R, respectively (Table 2). ‘Community Interest’ according to the Habitats Directive were identified through visual census. They belong to Macrophyta, Cnidaria, Bryozoa and Tunicata. Moreover, Census of species the analysis of ROV samples collected by grabber allowed identification of 12 Mediterranean endemic bryo- A total of 269 taxa (Appendix S1) were identified zoan species living on red algae which form maërl and/or through ROV videos, among which 185 were at species rhodoliths (Table 4). level. These taxa belong to: Macrophyta (Rhodophyta, Chlorophyta, Ochrophyta and Tracheophyta), Porifera, Cnidaria, Annelida, Mollusca, Bryozoa, Arthropoda, Graham Bank Echinodermata, and Chordata (Tunicata and Pisces). The number of taxa for the most abundant taxonomic groups The Ferdinandea shoal is the only investigated edifice is shown in Fig. 6. The largest number of taxa detected of this bank, falling within the euphotic zone (Fig. 2J). belonged to Porifera, Cnidaria and Pisces, with marked From the peak down to about 20/30 m this shoal shows a differences in identification level. Indeed, even though it highly complex, mainly rocky bottom, covered by dense

Medit. Mar. Sci., 22/2 2021, 297-316 301

http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 06/10/2021 09:44:35 | - Distribution of the main habitats along the ROV tracks on the high-resolution morpho-bathymetric map of the Graham Bank: A) volcanic cones; B) morpho-bathymetric map of B-E cones; B) morpho-bathymetric A) volcanic map of the Graham Bank: morpho-bathymetric tracks on the high-resolution along the ROV of the main habitats Distribution Fig. 2 : on point grabber sample and ROV habitats and semi-dark caves of maërl/rhodoliths D) distribution on B-E cones; habitats bottom detritic and coastal of coralligenous C) distribution cones; on F-I cones; H) distribution of corallige of bathyal muds habitat map of F-I cone; G) distribution on B-E cone; F) morpho-bathymetric B-E cones; E) distribution of shelf-edge rock habitat nous habitat on F-I cones; I) distribution of shelf-edge rock habitat on F-I cones; J) morpho-bathymetric map of J-M cones; K) distribution of coastal detritic bottom habitat on J-M cones; L) bottom habitat detritic of coastal map of J-M cones; K) distribution on F-I cones; J) morpho-bathymetric on F-I cones; I) distribution of shelf-edge rock habitat nous habitat map of N-Q on J-M cones; N) morpho-bathymetric and shelf-edge rock habitats algae of infralittoral on J-M cones; M) distribution habitats and maërl/rhodoliths of coralligenous, distribution rock habitat of shelf-edge point on N-Q cones; Q) distribution grabber sample and ROV habitat of maërl/rhodoliths on N-Q cones; P) distribution muds habitat of bathyal cones; O) distribution The scale bars of figures B-Q corresponds to 0.5 km. visual assessment. The habitats were identified through ROV on N-Q cones.

302 Medit. Mar. Sci., 22/2 2021, 297-316

http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 06/10/2021 09:44:35 | Fig. 3: Distribution of the main habitats along the ROV tracks on the high-resolution morpho-bathymetric map of the Nereo Bank: morpho-bathymetric map (A); distribution of infralittoral algae (B), coastal detritic bottom (C), coralligenous (D), maërl/ rhodoliths (E), and semi-dark caves (F) habitats; ROV grabber sample points (G). The habitats were identified through ROV visual assessment. populations of infralittoral algae, including Dictyota spp., was detected in the circalittoral disphotic zone associated Sargassum sp. and species belonging to the ‘Cystoseira’ with ‘coastal detritic’ bottoms. At 40-70 m depth, hard complex1. Between 30 and 40 m depth, an extensive and substrates host the ‘coralligenous biocoenosis’ (Fig. 2C, dense facies with species of Ceriantharia (Fig. 8A) was 2H, 2L) of two different aspects: one dominated by mac- spotted on a flat bottom composed of coarse volcanic rophytes (Sargassum spp., L. rodriguezii, and many other sand. This facies was found interspersed with ‘maërl-rh- unidentified brown and red algae) and one by megafauna odolith’ beds. All soft bottoms between 50 and 70/80 m (mainly gorgonians, such as Paramuricea clavata and Eu- depth on the Graham volcanic edifices host the ‘coastal nicella cavolinii, bryozoans, including Smittina cervicor- detritic bottom biocoenosis’, frequently characterised by nis, Hornera frondiculata, Margaretta cereoides, Myria- ‘maërl facies’ and by ‘association with rhodoliths’ (Fig. pora truncata, Pentapora fascialis, Reteporella spp., and 2C-D and 2K-L), mostly formed by P. calcareum and L. sponges such as Axinella polypoides and A. damicornis). corallioides (Table 3). Laminaria rodriguezii, an endem- The ‘association with Sargassum spp.’, the ‘facies with ic Mediterranean brown algae of conservation concern, E. cavolinii’, the ‘facies with P. clavata’ and ‘corallig-

1 The species previously allocated within the genus Cystoseira have been largely revised and new synonyms and allocations have been proposed (Orellana et al., 2019; Molinari & Guiry, 2020). Since identification at a specific level is not possible through video analysis, we indicate them with the term ‘Cystoseira’ complex.

Medit. Mar. Sci., 22/2 2021, 297-316 303

http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 06/10/2021 09:44:35 | Fig. 4: Distribution of the main habitats along the ROV tracks on the high-resolution morpho-bathymetric map of the Pantelleria Vecchia Bank: Α) morpho-bathymetric map; Β) distribution of coastal detritic bottom, C) infralittoral algae, D) coralligenous, E) Posidonia beds, F) maërl/rhodoliths, and G) semi-dark caves habitats. The habitats were identified through ROV visual assessment. enous platforms’ were visually distinguished. At this um rubrum (Fig. 8C), Leiopathes glaberrima (Fig. 8D), bathymetric range, transitional areas (ecotones) between Dendrophyllia ramea (Fig. 8E), Savalia savaglia (Fig. ‘coastal detritic bottom’ and ‘coralligenous biocoenosis’ 8F), Callogorgia verticillata and Parantipathes larix. are also conspicuous, characterised by soft corals such Most of these species are protected by international direc- as Paralcyonium spinulosum, rhodoliths and soft-bodied tives and conventions and are included in the threatened algae such as L. rodriguezii. The ‘semi-dark cave bioco- categories of the IUCN Anthozoan Red List (Table 3). enosis’ is also present in overhangs in both infralittoral These corals can form dense monospecific gardens (C. and circalittoral zones (Fig. 2D). This biocoenosis shows rubrum) and/or multi-specific gardens (anthipatharians different aspects depending on the occurrence and domi- and gorgonians), hosting epibenthic Echinodermata such nance of species, mainly encrusting Porifera and Cnidar- as Astrospartus mediterraneus. It is worth noting that the ia such as Hexadella racovitzai, Leptopsammia pruvoti observed facies with C. rubrum was not the one occur- and Parazoanthus axinellae. These species were often ring within the ‘semi-dark cave biocoenosis’, listed in the grouped in almost monospecific aggregations, such as the SPA/BD Protocol as a priority habitat (IV.3.2.2.), but a ‘facies with P. axinellae’. Between 100 and 200 m, the facies neither described nor listed in the SPA/BD Proto- ‘shelf-edge rock biocoenosis’ is present with flourishing col, which develops on open rocks and on gently sloping coral gardens (Fig. 2E, 2I, 2M, 2Q and Fig. 8B) com- substrates and is characterised by upwardly growing red posed of black corals, zoanthids and gorgonians, such as coral colonies. Moreover, on the Graham Bank and in its Antipathes dichotoma, Antipathella subpinnata, Coralli- vicinity alongside some volcanic reliefs, at 140-180 m

304 Medit. Mar. Sci., 22/2 2021, 297-316

http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 06/10/2021 09:44:35 | Table 2. Habitat types (including benthic biocoenoses, with relative associations and/or facies) detected on the Graham, Nereo and Pantelleria Vecchia Banks by ROV video recordings. The habitat classification‡ follows the identification sheets of priority habitats according to the SPA/BD Protocol (Relini & Giaccone, 2009) and reports the SPA/BD Protocol (Barcelona Convention) codes and the correspondent Habitats Directive (92/43/EEC) identification codes (Annex I ‘Natural habitat types of Community Interest whose conservation requires the designation of Special Areas of Conservation’; EU, 1992).

Biocoenosis Association A./Facies F. Habitat Dir. SPA/BD Protocol Bank† Range depth Identification Identification code (m) code (Classification) ‡ Posidonia oceanica A. Posidonia oceanica 1120 III.5.1 (P) Pa 20-40 beds Biocoenosis of coarse A. with rhodoliths - III.2.2.2. (P) Gr, Ne, Pa 90-120 (Gr) sands and fine gravels 30-50 (Ne, Pa) under the influence of bottom currents

Biocoenosis of coastal F. maërl - III.3.2.1. (P) Gr, Ne, Pa 90-120 (Gr) detritic bottom 30-50 (Ne, Pa) A. Laminaria rodriguezii - IV.2.2.7. (P) Gr, Ne 40-60

A. with rhodoliths - IV.2.2.1. (OH) Gr, Ne, Pa 80-100 (Gr) 50-55 (Ne, Pa) F. maërl - IV.2.2.2. (R) Gr, Ne, Pa 80-100 (Gr) 50-55 (Ne, Pa) Biocoenosis of - 1170 III.6.1. (P) Gr, Ne, Pa 40-60 (Gr) infralittoral algae 30-50 (Ne, Pa) A./F. Coralligenous in enclaves 1170 III.6.1.35. (P) Pa 30-50

Biocoenosis of - 1170 IV.3.1. (P) Gr, Ne, Pa 25-50 coralligenous A. Sargassum spp. 1170 IV.3.1.5. (P) Gr, Pa 20 F. Eunicella cavolinii 1170 IV.3.1.10. (P) Gr 50-100

F. Eunicella singularis 1170 IV.3.1.11. (P) Ne 30-55

F. Paramuricea clavata 1170 IV.3.1.13. (P) Gr 50-120

Coralligenous platforms 1170 IV.3.1.15. (P) Gr 80-160

Biocoenosis - 1170 IV.3.3. (R) Gr 80-160 of shelf-edge rock Biocoenosis of semi- - 8330 IV.3.2. (P) Gr, Ne, Pa 80 (Gr) dark caves 50 (Ne) 20-40 (Pa) F. Parazoanthus axinellae 8330 IV.3.2. 1. (R) Gr, Ne, Pa 80

Biocoenosis of - - V.1.1. (R) Gr 180 bathyal muds F. soft muds with Funiculina - V.1.1.3. (P) Gr 180 quadrangularis

Medit. Mar. Sci., 22/2 2021, 297-316 305

http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 06/10/2021 09:44:35 | Fig. 5: Number of identified habitats for each degree (low, Fig. 6: Number of taxa identified at species level or higher tax- medium, high) assigned to the following habitat variables: vul- onomic rank including genus, family and even phylum level nerability, rarity, heritage value, aesthetic value and economic through ROV visual assessment on the Graham, Nereo and Pan- value, following the classification proposed by Relini & Giac- telleria Vecchia banks. cone (2009).

Fig. 7: Number of species detected on the Graham, Nereo and Pantelleria Vecchia banks through ROV visual assessment included in the threatened categories of the Global, Mediterranean and/or Italian IUCN Red Lists (IUCN, 2015; Italian Committee of IUCN, 2016). The total number of species in each category is shown (dark triangles); note that species can be present in more than one list. depth on the muddy sea floor, three areas were observed, extensive facies with F. quadrangularis was observed on including dead fragments (thanatocoenoses) of red coral ‘bathyal muds’ (Fig. 8H). Between 100 and 150 m, evi- and Dendrophyllia spp. According to Lodolo et al. (2017) dence of hydrothermal vents was detected, consisting of these deposits have accumulated primarily (or partly) as gas emissions associated with white and orange mineral- a consequence of volcanic activity dislodging living (or isations. dead) corals from the steep flanks of the volcanoes on which they lived. The large hydroid Lytocarpia myriophyllum and the Nereo Bank sea pen Virgularia mirabilis form dense aggregations on detritic bottoms at 140-180 m depth. At about 130 m, The euphotic-to-disphotic bottoms between 30 and the whip coral Viminella flagellum is abundant, forming 45 m are dominated by macrophytes. Rocky and soft dense monospecific forests on rocky substrates covered substrata are covered by extensive and diversified algal with a thin mud veneer (Fig. 8G). At about 190 m, a quite populations. A dense population of the kelp L. rodriguezii

306 Medit. Mar. Sci., 22/2 2021, 297-316

http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 06/10/2021 09:44:35 | Table 3. Species of conservation concern, detected during ‘ORBS Banchi 2014-2015’ research surveys, whose conservation or exploitation is regulated by international Directives and Conventions: CITES‡, BERN§, HABITAT¶, SPA/BD††. The species included in the Global, Mediterranean and/or Italian IUCN Red Lists of Threatened Species are also reported. In the last column the Population Trend at Global1, Mediterranean2 and/or Italian3 (International Union for Conservation of Nature, 2015; Italian Committee of International Union for Conservation of Nature, 2016) scale was shown.

Directive/ IUCN IUCN IUCN Population TAXA Convention Global1 Mediterranean2 Italian3 Trend Arthropoda Pachylasmus giganteum P2, B2 - - - - Palinurus elephas B3, P3 - - - - Cnidaria Acanthogorgia hirsuta - - LC LC unknown2,3 Alcyonium acaule - - LC DD decreasing2 Alcyonium coralloides - - LC LC stable2,3 Alcyonium palmatum - - LC LC decreasing2; unknown3 Alicia mirabilis - - LC LC unknown2; stable3 Amphianthus dohrnii - - DD - unknown2,3 Anemonia viridis - - LC LC stable2,3 Antipathella subpinnata CB, B3, P2 - NT LC decreasing2 Antipathes dichotoma CB, B3, P2, P3 - NT LC decreasing2 Astroides calycularis B2, P2, CB - LC LC stable2 Bebryce mollis - - DD DD unknown2,3 Callogorgia verticillata P2 - NT NT decreasing3 Caryophyllia cyathus CB - DD DD unknown2,3 Cereus pedunculatus - - LC LC stable2,3 Cerianthus membranaceus - - LC LC unknown2; stable3 Cladocora caespitosa CB, P2 E E LC decreasing1,2; stable3 Cladopsammia rolandi - DD DD unknown2,3 Condylactis aurantiaca - LC LC DD unknown1,2,3 Corallium rubrum B3, H5, P3 - E E decreasing2,3 Corynactis viridis - - LC LC stable23 Cribrinopsis crassa - - LC DD unknown2,3 Dendrophyllia cornigera P2, CB - E V decreasing2,3 Dendrophyllia ramea P2, CB - V DD unknown2,3 Eunicella cavolinii - - - LC stable3 Eunicella singularis - - NT V decreasing2,3 Funiculina quadrangularis - - V CE decreasing2,3 Leiopathes glaberrima CB, B3, P2 - E E decreasing2,3 Leptopsammia pruvoti CB - LC LC stable3 Lytocarpia myriophyllum - - LC - unknown2 Paracyathus pulchellus - - DD DD unknown2,3 Paralcyonium spinulosum - - LC LC stable3 Paramuricea clavata - - V LC decreasing2 Paramuricea macrospina - - DD LC unknown2,3 Parantipathes larix CB, P2 - NT LC decreasing2 Parazoanthus axinellae - - LC LC stable2,3 Pennatula rubra - - V DD decreasing2; unknown3 Phymanthus pulcher - - DD DD unknown2,3 Pteroeides griseum - - V DD decreasing2; unknown3 continued

Medit. Mar. Sci., 22/2 2021, 297-316 307

http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 06/10/2021 09:44:35 | Table 3 continued Directive/ IUCN IUCN IUCN Population TAXA Convention Global1 Mediterranean2 Italian3 Trend Savalia Savaglia B2, P2 - NT NT decreasing2,3 Swiftia pallida - - DD DD unknown2,3 Thalamophyllia gasti - - DD DD unknown2,3 Veretillum cynomorium - - LC DD unknown2,3 Villogorgia bebrycoides - - DD DD unknown2,3 Viminella flagellum - - NT LC unknown2,3 Virgularia mirabilis - - LC V unknown2; decreasing3 Echinodermata Centrostephanus longispinus H4, P2, B2 - - - - Paracentrotus lividus P3, B3 - - - - Macrophyta - “Cystoseira” complex B1, P2 - - - - Laminaria rodriguezii B1, P2 - - - - Lithothamnion corallioides‡‡ H5 - - - - Phymatolithon calcareum ‡‡ H5 - - - - Posidonia oceanica B1, P2 LC LC - decreasing1,2 Sargassum spp. P2 - - - - Mollusca Octopus vulgaris - LC LC - unknown1,2 Pisces Anthias anthias - LC LC LC stable1,2,3 Apogon imberbis - LC LC LC stable1,2,3 Aulopus filamentosus - LC LC LC unknown1,2; stable3 Boops boops - LC LC LC stable1,2,3 Callanthias ruber - LC LC LC unknown1,2; stable3 Capros aper - LC LC LC unknown1,2; increasing 3 Chromis chromis - LC LC LC stable1,2,3 Conger conger - LC LC LC increasing1; stable3 Coris julis - LC LC LC stable1,3 Ctenolabrus rupestris - LC LC LC stable1,2,3 Dasyatis pastinaca - DD V DD decreasing2 Diplodus vulgaris - LC LC LC stable2,3 Epinephelus caninus - DD DD DD unknown1,2,3 Epinephelus costae - DD DD V unknown1,2; decreasing3 Epinephelus marginatus P3, B3 V E E decreasing1,2; increasing3 Gadella maraldi - LC LC LC unknown1,2,3 Helicolenus dactylopterus - LC LC LC unknown1,2,3 Hyporthodus haifensis - DD DD DD unknown1,2,3 Labrus merula - LC LC LC stable1,2,3 Labrus mixtus - LC LC LC stable1,2,3 Labrus viridis - V V LC decreasing1,2; stable3 Lappanella fasciata - LC - LC stable1,3 Macroramphosus scolopax - LC LC LC unknown1,2; stable3 Myliobatis aquila - DD V LC decreasing2,3 Mullus surmuletus - LC LC LC stable2,3 Muraena helena - LC LC LC unknown1,2; increasing3

continued 308 Medit. Mar. Sci., 22/2 2021, 297-316

http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 06/10/2021 09:44:35 | Table 3 continued Directive/ IUCN IUCN IUCN Population TAXA Convention Global1 Mediterranean2 Italian3 Trend Pagellus bogaraveo - NT LC LC decreasing1,3; increasing3 Parablennius rouxi - LC LC LC stable1,2,3 Phycis phycis - LC LC LC unknown1,2; increasing3 Scorpaena elongata - LC LC LC unknown1,2; stable3 Scorpaena maderensis - LC LC LC unknown1,2; stable3 Scorpaena notata - LC LC LC unknown1,2; stable3 Scorpaenodes arenai - DD DD LC unknown1,2; stable3 Seriola dumerili - LC LC LC stable2,3 Serranus cabrilla - LC LC LC stable1,2,3 Serranus scriba - LC LC LC stable1,2,3 Sparus aurata - LC LC LC stable1,2,3 Spicara maena - LC LC LC decreasing2; stable3 Spicara smaris - LC LC LC decreasing2; stable3 Spondyliosoma cantharus - LC LC LC stable2,3 Symphodus doderleini - LC LC LC stable1,2,3 Symphodus mediterraneus - LC LC LC stable1,2,3 Symphodus melanocercus - LC LC LC stable1,2,3 Symphodus ocellatus - LC LC LC stable1,2,3 Symphodus roissali - LC LC LC stable1,2,3 Symphodus tinca - LC LC LC stable1,2,3 Thalassoma pavo - LC LC LC stable1,2,3 Trachurus trachurus - V LC LC stable1,2,3 Zeus faber - DD LC LC stable1,3 Porifera Axinella damicornis - - - LC unknown3 Axinella polypoides B2, P2 - - E unknown3 Agelas oroides - - - LC unknown3 Dysidea avara - - - LC unknown3 Pachastrella monilifera - - - V unknown3 Chondrosia reniformis - - - LC unknown3 Cliona celata - - - LC unknown3 Ircinia variabilis - - - LC unknown3 Petrosia ficiformis - - - V unknown3 Poecillastra compressa - - - V unknown3 Spongia lamella B3, P3 - - E unknown3

‡CB: CITES, Convention on International Trade in Endangered Species of Wild Fauna and Flora (C). Annex: B-Species that are not necessarily now threatened with extinction but that may become so unless trade is closely controlled. §B1, B2, B3: BERN, “Convention on the Conservation of European Wildlife and Natural Habitats(B). Annex: 1-Strictly protected flora species; 2-Strictly protected fauna species; 3-Protected fauna species. ¶H4, H5: HABITAT 42/93 CEE Directive (H). Annex: 4-Animal and plant species of community interest in need of strict protection; 5-Animal and plant species of community interest whose taking in the wild and exploitation may be subject to management measures. ††P2, P3: SPA/BD, Protocol Concerning Specially Protected Areas and Biological Diversity of the Barcelona Convention (P). Annex: 2-List of endangered or threatened species; 3-List of species whose exploitation is regulated IUCN: CE: Critically Endangered; E: Endangered; V: Vulnerable; NT: Near threatened; LC: Least Concern; DD: Data Deficient.

Medit. Mar. Sci., 22/2 2021, 297-316 309

http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 06/10/2021 09:44:35 | (Fig. 9A) is widespread, accompanied by other brown al- Eunicella singularis and the candelabrum-shaped sponge gae such as Dictyopteris cfr. polypodioides, Dictyota spp., Raspailia viminalis (Fig. 9G). In restricted infralittoral Sargassum sp. and species of the ‘Cystoseira’ complex, as and circalittoral zones, the ‘semi-dark caves biocoenosis’ well as by red algae such as Lophocladia lallemandii and (Fig. 3F) was recognised as well. At about 50 m depth, a Osmundaria volubilis. At 30-60 m depth, gently sloping school of more than 30 individuals of the common eagle rocky bottoms host the ‘coralligenous biocoenosis’ (Fig. ray Myliobatis aquila was observed (Fig. 9H). 3D). At about 30 m, the ‘facies with Eunicella singularis’ is widespread, characterised by the abundance of this gorgonian in flourishing macrophyte communities (Fig. 9C). Pantelleria Vecchia Bank At 40-50 m, a highly diversified ‘coralligenous biocoenosis’ is frequently spotted on vertical rocky walls, dom- The euphotic bottoms are dominated by flourishing P. inated by the encrusting sponge Hexadella racovitzai, oceanica seagrasses (Fig. 4E and Fig. 10A) on both rocky the tubular sponge Haliclona sp., and the erect ascidians and soft bottoms, interspersed with diversified algal pop- Aplidium cfr. proliferum and Aplidium cfr. tabarquensis ulations, mainly of Sargassum sp. (Fig. 10B), Flabellia (Fig. 9D). These rocky habitats offer numerous refuges petiolata, Osmundaria volubilis, and species of the ‘Cys- to fish and decapod crustaceans of high commercial val- toseira’ complex, which also thrive on substrates of the ue. Epinephelus marginatus, assessed as an endangered disphotic zone. The ‘biocoenosis of semi-dark caves’ is species in the Global and Mediterranean IUCN Red Lists present, mainly on the sciaphilous subvertical hard sub- (IUCN, 2015; Italian Committee of IUCN, 2016; Table 3) strates, as well as in enclaves of the infralittoral zone (Fig. has been observed, together with Palinurus elephas (Fig. 4G). As on the other banks, the ‘facies with P. axinellae’ 9E), whose exploitation is regulated (Table 3). At 30-60 m was detected. Extensive ecotones between the ‘associa- depth, on soft bottoms the ‘coastal detritic bottom bioco- tion with rhodoliths’ and ‘coralligenous’ were detected, enosis’ (Fig. 3C) is represented by the ‘maërl facies’ and chiefly around bioconstructions (Fig. 4D, 4F and Fig. the ‘association with rhodoliths’ (Table 3 and Fig. 3E). 10C). The ‘coralligenous’ was detected mainly on ver- Locally dense populations of the soft coral Veretillum tical rocky walls but also in enclaves in the infralittoral cynomorium (Fig. 9F) occur. In this bathymetric range, zone. This habitat is markedly diversified, dominated by ecotones between ‘coastal detritic bottom’ biocoenosis different species, among which are Leptopsammia pru- and ‘coralligenous’ (Fig. 3D), bioconstructions are also voti and Astroides calycularis (Fig. 10D–E). On the Pan- conspicuous on this bank. Rocks frequently host sponge– telleria Vecchia Bank, the group of protected species or cnidarian assemblages dominated by Alcyonium acaule, whose exploitation is regulated is represented by Posido- nia oceanica, Astroides calycularis, Axinella polypoides, Table 4. Mediterranean endemic species detected on the Gra- Leptopsammia pruvoti and Centrostephanus longispinus ham, Nereo and Pantelleria Vecchia banks through ROV visual (Fig. 10F). The endangered dusky grouper Epinephelus assessment and ROV grabber sample †. marginatus (Fig. 10G) and the vulnerable common eagle ray Myliobatis aquila were also detected. Taxonomic group Mediterranean endemic species

Macrophytes Laminaria rodriguezii Discussion Posidonia oceanica The present study evaluated, for the first time, the bio- Cnidaria Cladopsammia rolandi diversity of the Graham, Nereo and Pantelleria Vecchia Condylactis aurantiaca Banks located in the Strait of Sicily. The numerous banks Eunicella cavolinii in the area, and most of the shallower ones located in the euphotic zone (0-100 m), provide habitats for a large va- Bryozoa Adeonella calveti riety of species together with unique ecosystems, many Bertorsonidra prenanti† of which are still to be discovered and described (Clark Cellepora adriatica† et al., 2006), as demonstrated by the newly reported Cla- Coronellina fagei† docora coespitosa–Treptacantha ballesterosii habitat Distansescharella seguenzai† (Pons-Fita et al., 2020). This also applies to the ‘Coralli- Gregarinidra gregaria† um rubrum facies’ observed on the Graham Bank, which Hippopleurifera pulchra† would need a formal description prior to its possible in- ‘Microecia’ occulta† clusion in lists of marine habitats. Patinella mediterranea† The use of ROV technology on the Graham, Nereo Rhynchozoon pseudodigitatum† and Pantelleria Vecchia Banks allowed the recording of Stephanoteca arrogata† an unexpectedly high richness of species (identified by Stephanoteca monoecensis† taxonomists at different levels: species, genus, family, Tubulipora hemiphragmata† phylum) and habitats of conservation concern otherwise unattainable by traditional sampling methods. In fact, Tunicata Halocynthia papillosa ROVs equipped with high-definition digital cameras are able to operate in natural and artificial complex habitats,

310 Medit. Mar. Sci., 22/2 2021, 297-316

http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 06/10/2021 09:44:35 | Fig. 8: ROV images of some habitats and species identified on the Graham Bank. A) Dense population of Ceriantharia sp. on a gently sloping detritic bottom (Ferdinandea Bank: –30 m); B) Deep coral garden (–140 m), dominated by the gorgonians Eunicella cavolinii and Paramuricea clavata, hosting different schools of fish, among which are Macroramphosus scolopax and Callanthias ruber; C) A dense monospecific deep coral garden (–110 m) composed of Corallium rubrum hosting the ophiuroid Astrospartus mediterraneus; D) A large colony of the antipatharian Leiopathes glaberrima on a slightly silted rocky substrate (–140 m); E) Col- ony of the scleractinian Dendrophyllia ramea (–100 m); F) colony of Savalia savaglia (–80 m); G) Deep megabenthic community thriving on a highly silted hardground covered by a Viminella flagellum forest, massive sponges and the alcyonacean Chirone- phthya mediterranea. Scattered colonies of Bebryce mollis and Eunicella cavolinii are also visible; H) A specimen of the tall sea pen, Funiculina quadrangularis, on mud (–190 m). Scale bars 10 cm. even at depths where diving is logistically limited (Con- ROV explorations also allowed the observation of soli et al., 2016, 2018); moreover, being non-invasive, some rare or poorly known fish species, such as Scor- they can also be used to investigate sensitive habitats paenodes arenai, Gadella maraldi, Hyporthodus haifen- (Consoli et al., 2015, 2018; Sinopoli et al., 2015). sis and Myliobatis aquila (Battaglia et al., 2015). Also, Using this technological approach, an extensive num- the occurrence of the whip-like gorgonian Viminella fla- ber of associations and facies of conservation concern gellum is worthy of note because it forms dense monospe- were observed on all three banks, mainly on rocky sub- cific forests or ‘deep coral gardens’ on the Graham Bank. strates that dominate from a few metres down to 250 m This species, previously known from near the Gibraltar depth. In the studied area, the occurrence of hard bot- Strait and the Balearic Sea, has recently been reported in toms, often of volcanic origin, and also at depths usually rocky habitats at 100-250 m depth from the Ligurian Sea covered with soft substrates, promotes a diversification to the Strait of Sicily (Angiolillo et al., 2014). and richness of habitats and species in the area. The Graham Bank hosts the largest variety of habitats

Medit. Mar. Sci., 22/2 2021, 297-316 311

http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 06/10/2021 09:44:35 | Fig. 9: ROV images of some habitats and species identified on the Nereo Bank. A) Dense algal population dominated by Lami- naria rodriguezii (–40 m); B) Gently sloping coralligenous bottom composed of sponges such as Cliona celata, cnidarians such as Alcyonium acaule, and numerous ascidians and bryozoans (–50 m); C) Gently sloping coralligenous bottom dominated by Euni- cella singularis scattered in a dense algal population (–30 m); D) Highly diversified vertical coralligenous community dominated by the encrusting sponge Hexadella racovitzai, the tubular Haliclona sp. and the erect ascidians Aplidium cfr. proliferum and Aplidium cfr. tabarquensis (–40 m); E) A specimen of Palinurus elephas emerging from a coralligenous concretion (–50 m); F) Coastal detritic bottom dominated by the finger-shaped sea pen Veretillum cynomorium (–50 m); G) Sponge–cnidarian assemblage dominated by Alcyonium acaule, Eunicella singularis and the candelabrum-shaped sponge Raspailia viminalis (–50–60 m); H) Specimens of Myliobatis aquila (–40 m). Scale bars 10 cm. and species of conservation concern, likely to be due to or abandoned fishing gear, represents a serious threat to its wider bathymetric range from the infralittoral to the bank ecosystems by entangling and damaging the sessile upper bathyal zone. The high RLI value indicated that fauna, especially corals (Octocorallia and Hexacorallia; the Cnidaria and Pisces in these biocoenoses are gen- Consoli et al., 2018). erally not in ‘danger’ of extinction. Nevertheless, most The results of this study provide a fundamental base- species found in the ‘shelf-edge rock’ and in the ‘bathyal line for addressing more in-depth studies on macro- mud’ habitats are included in the Threatened categories phytes, which appear to be very abundant and diversi- in the IUCN Red List: they were represented by cni- fied on these banks. In particular, further research should darians such as gorgonians, antipatharians and pennatu- be focused on the species belonging to the ‘Cystoseira’ laceans. Moreover, many of the cnidarian species found complex, and Sargassum sp., Laminaria rodriguezii and in bank areas and for which data about conservation state Posidonia oceanica populations. Porifera need further in- are available (Global, Mediterranean and/or Italian Red vestigation as well, owing to their high diversity, mainly Lists), are Near Threatened with extinction and show a in coralligenous and semi-dark cave habitats. Numerous decreasing Population Trend. Furthermore, the abundant specimens of Pachastrella monilifera and Poecillastra marine litter present in the area, composed mostly of lost compressa were found on the Graham Bank between 100

312 Medit. Mar. Sci., 22/2 2021, 297-316

http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 06/10/2021 09:44:35 | Fig. 10: ROV images of some habitats and species identified on the Pantelleria Vecchia bank. A) Posidonia oceanica seagrass flourishing on rocky outcrops (–20 m); B) Algal population dominated by Sargassum sp. densely covering the euphotic bottoms along with Posidonia oceanica seagrass (–20–30 m); C) Coastal detritic bottom with rhodoliths and cidarid sea urchins (–50 m); D) Vertical surface hosting a coralligenous community dominated by Astroides calycularis. Wide patches of the encrusting sponges Hexadella racovitzai are also visible (–35 m). E) Vertical-to-overhanging surface hosting a coralligenous community dominated by Leptopsammia pruvoti (–35 m); F) The protected urchin Centrostephanus longispinus (–30 m); G) Two adults of the endangered dusky grouper Epinephelus marginatus (–15 m). Scale bars 10 cm. and 180 metres depth on rocky substrates. A recent study in similar habitats and with comparable high densities (Bo et al., 2012b) reported the first record of the so-called (Marzia Bo, personal observation). deep sponge grounds, composed mainly of these two as- The IUCN Red Lists and the international conven- trophorids, on some seamounts in the Mediterranean Sea. tions and directives represented a baseline for this study; It would be interesting to investigate the occurrence of however, despite the different lists of protected species these sponge assemblages on the Graham Bank. More- consulted, it was possible to make only limited consid- over, it would be important to elucidate better the large erations on the state of conservation of the identified diversity of the banks’ sponges from the taxonomic point species. In fact, some taxonomic groups, such as Mac- of view, as numerous other un-identifiable sponge species rophyta, Bryozoa, Tunicata and Porifera are scarcely or were found to form important aggregations. Cnidarians not considered in these lists. This could be for different such as Chironephthya cfr. mediterranea, Aiptasia sp., reasons, such as the low commercial value, the micro- and Thalamophyllia gasti deserve further analysis, due scopic dimensions (especially Bryozoa) and the difficulty to their rarity and their ability to contribute to the struc- of collection and identification. tural complexity of the coral gardens. Dense population As stated by Moser et al. (2016), remarkable discrep- of Ceriantharia sp. (probably Arachnantus oligopodus) ancies in species conservation status assessment exist observed on the Graham Bank need a deep taxonomic between the IUCN Red Lists and the international con- review. This species, an enigmatic cerianthid, was recent- ventions and directives (Habitats Directive, Bern Con- ly found in the Ligurian Sea and Central Tyrrhenian Sea vention, SPA/BD Protocol, CITES). This was confirmed

Medit. Mar. Sci., 22/2 2021, 297-316 313

http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 06/10/2021 09:44:35 | by our results; in fact, out of 115 taxa of conservation mended in the Annex I of SPA/BD Protocol, reporting concern observed on the banks of the Strait of Sicily, 108 the criteria for the choice of protected marine and coastal are listed in the IUCN lists, 22 in the SPA/BD protocols, areas that could be included in the SPAMI list. 16 in the Bern Convention, 10 in the CITES and only four are included in the Habitats Directive. Moreover, very few species that are threatened with extinction or Acknowledgements are endangered or vulnerable according to the IUCN Red Lists are listed in the Habitat Directive. This paper is dedicated to Giovanni Bortoluzzi who Not surprisingly, the IUCN Red Lists, although not attended, with passion, commitment and professional- legally binding for European States (unlike the Direc- ity, to the ‘ORBS Banchi 2014-2015’ research surveys, tives and Conventions), represent the most important and who unexpectedly passed away in October 2015. instruments for evaluating the extinction risk of species We thank Pietro Battaglia, Silvana Campagnuolo, Luca worldwide (Moser et al., 2016); they also contain most Castriota, Manuela Falautano, and Mauro Sinopoli for complete lists of species. In spite of this, the conservation participating to the ‘ORBS Banchi 2014-2015’ research policies in the EU are largely focused on the Habitats Di- surveys. Alfio Viola (University of Catania) is acknowl- rective, which is really not exhaustive. edged for SEM assistance. Paper financially supported by It follows that at European scale an exhaustive and Linea di intervento 3.2.1.2 of POR FESR Sicilia 2007- legally binding instrument to protect species of conserva- 2013 aimed to assess the marine biodiversity in Sicily. tion concern is still lacking, and its preparation, approval Additional funds were provided by the University of and issue are highly recommended as the starting point to Catania through “Piano per la Ricerca 2016-2018” to A. suggest measures to be undertaken for their protection. Rosso, n. 22722132118 and “PiaCeRi-Piano Incentivi per The ecological and biological data here reported la Ricerca di Ateneo 2020-22 linea di intervento 2”. This are complementary to those reported by Altobelli et al. is the contribution n. 472 of the Catania Palaeontological (2017); these authors carried out an ecological evalua- Research Group. tion of the Mediterranean banks. Data acquired to date allow to state that these three banks are of high ecological and biological value. However, they are endangered by References human activities: mainly fishing, which presently represents the main source of marine litter, and oil and gas Alongi, G., Cormaci, M., Furnari, G., 2002. The Corallinaceae extraction, whose increase is foreseen in the near future. (Rhodophyta) from the Ross Sea (Antarctica): a taxonomic In general, research has shown the uniqueness of these revision rejects all records except Phymatolithon foecun- offshore shallow environments, which can represent dum. Phycologia, 41 (2), 140-146. a network of biodiversity hot spots at a Mediterranean Altobelli, C., Perzia, P., Falautano, M., Consoli, P., Canese, S. scale, that should be managed through binding, region- et al., 2017. Mediterranean banks in EBSA area: Hotspots ally based and supranational collaborative management of biodiversity under threat. Marine Environmental Re- measures (Altobelli et al., 2017; Consoli et al., 2018). search, 131, 57-68. The whole Strait of Sicily has been recently consid- Angiolillo, M., Bavestrello, G., Bo, M., Cau, A., Cau, A. et al., ered as a relevant area of conservation concern (CBD- 2014. Distribution of the deep-dwelling gorgonian Viminel- COP 12 Decisions, 2014). In fact, it has been included in la flagellum in the Italian western Mediterranean Sea by the 12 priority conservation areas likely to contain candi- means of multi-year ROV surveys, Vol. 31. p. 65-66. In: date sites for SPAMIs and included in the EBSA by the Proceedings of the 1st Mediterranean symposium on the Contracting Parties of the Convention on Biological Di- conservation of Dark Habitats, Portorož, Slovenia, 31 Oct versity for their high naturalistic importance. The results 2014. RAC/SPA publications, Tunis. acquired could be useful for the delimitation of a future Angiolillo, M., di Lorenzo, B., Farcomeni, A., Bo, M., Baves- SPAMI or a network of MPAs (including the investigated trello, G. et al., 2015. Distribution and assessment of ma- banks and others located in the same area) and the iden- rine debris in the deep Tyrrhenian Sea (NW Mediterranean tification of zones within them with different area-based Sea, Italy). Marine Pollution Bulletin, 92 (1-2), 149-159. management measures. An example of this approach is Ban, N.C., Bax, N.J., Gjerde, K.M., Devillers, R., Dunn, D.C. suggested by the Man and the Biosphere (MAB) Pro- et al., 2014. Systematic conservation planning: a better rec- gramme for the designation of UNESCO biosphere re- ipe for managing the high seas for biodiversity conservation serves (UNESCO, 2013). In such reserves the core area and sustainable use. Conservation Letters, 7 (1), 41-54. is devoted to long-term protection with specific conser- Battaglia, P., Canese, S., Ammendolia, G., Romeo, T., Sandulli, vation objectives; in the buffer zone only activities com- R. et al., 2015. New records and underwater observation patible with the conservation objectives can take place; of the rare fish Scorpaenodes arenai (Osteichthyes: Scor- in the transition area sustainable resource management paenidae) from the central and western Mediterranean Sea. practices are promoted and developed. Italian Journal of Zoology, 82 (3), 454-458. In order to protect these banks, the future challenges Bo, M., Canese, S., Spaggiari, C., Pusceddu, A., Bertolino, M. are to promote further scientific research, strengthen in- et al., 2012a. Deep coral oases in the south Tyrrhenian sea. ternational political collaboration and foster the involve- PLoS ONE, 7 (11), e49870. ment of local stakeholders and populations, as recom- Bo, M., Bertolino, M., Bavestrello, G., Canese, S., Giusti, M.

314 Medit. Mar. Sci., 22/2 2021, 297-316

http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 06/10/2021 09:44:35 | et al., 2012b. Role of deep sponge grounds in the Mediter- for Signature on 19 September 1979. Council of Europe, ranean Sea: a case study in southern Italy. Hydrobiologia, European Treaty Series, No. 104, 11 pp. 687 (1), 163-177. Danovaro, R., Gambi, C., Dell’Anno, A., Corinaldesi, C., Fra- Bo, M., Bava, S., Canese, S., Angiolillo, M., Cattaneo-Vietti, R. schetti, S. et al., 2008. Exponential decline of deep-sea et al., 2014. Fishing impact on deep Mediterranean rocky ecosystem functioning linked to benthic biodiversity loss. habitats as revealed by ROV investigation. Biological Con- Current Biology, 18 (1), 1-8. servation, 171, 167-176. EU, 1992. Directive 92/43/EEC of Council of the European Boyes, S.J., Elliott, M., 2014. Marine legislation - The ultimate Communities of 21 May 1992 on the conservation of natu- “horrendogram”: International law, European directives & ral habitats and of wild fauna and flora. Official Journal of national implementation. Marine Pollution Bulletin, 86 (1), the European Union, L206, 7-50. 39-47. EU, 2008. Directive 2008/56/EC (Marine Strategy Framework Butchart, S.H., Akçakaya, H.R., Chanson, J., Baillie, J.E., Directive) of the European Parliament and of the Council Collen, B. et al., 2007. Improvements to the red list index. of 17 June 2008 establishing a framework for community PLoS ONE, 2 (1), e140. action in the field of marine environmental policy. Official Cardinale, B.J., Duffy, J.E., Gonzalez, A., Hooper, D.U., Per- Journal of the European Union, L164, 19-40. rings, C. et al., 2012. Biodiversity loss and its impact on EU, 2009. Directive 2009/147/EC of the European Parliament humanity. Nature, 486 (7401), 59. and of the Council of 30 November 2009 on the conserva- Civile, D., Lodolo, E., Tortorici, L., Lanzafame, G., Brancolini, tion of wild birds. Official Journal of the European Com- G., 2008. Relationships between magmatism and tectonics munities, L20, 7-25. in a continental rift: the Pantelleria Island region (Sicily European Commission, 2013. Interpretation Manual of Euro- Channel, Italy). Marine Geology, 251 (1-2), 32-46. pean Union Habitats, version EUR 28. DG Environment, Clark, M.R., Tittensor, D., Rogers, A.D., Brewin, P., Schlacher, Nature ENV B.3, 144 pp. T. et al., 2006. Seamounts, deep-sea corals and fisheries: Falzone, G., Lanzafame, G., Rossi, P., 2009. L’isola che non vulnerability of deep-sea corals to fishing on seamounts be- c’è: il vulcano Ferdinandea nel Canale di Sicilia. Geoitalia, yond areas of national jurisdiction. UNEP-WCMC, Cam- 29, 15-21. bridge, UK, 86 pp. Fiorentino, F., Bono, G., Gancitano, V., Garofalo, G., Gristina, Colantoni, P., Cremona, G., Ligi, M., Borsetti, A.M., Cati, M. et al., 2011. Caratterizzazione ambientale delle aree di F., 1985. The Adventure Bank (off south-western Sicily): pesca - GSA 16 - Coste meridionali della Sicilia, p. 66-72. a present day example of carbonate shelf sedimentation. In: Lo stato della pesca e dell’acquacoltura nei mari ita- Giornale di Geologia, 47 (1), 165-180. liani. Cataudella, S., Spagnolo, M. (Eds). Ministero delle Coll, M., Piroddi, C., Steenbeek, J., Kaschner, K., Lasram, Politiche Agricole Alimentari e Forestali, Roma. F.B.R. et al., 2010. The biodiversity of the Mediterranean Hochkirch, A., Schmitt, T., Beninde, J., Hiery, M., Kinitz, T. Sea: estimates, patterns, and threats. PLoS ONE, 5 (8), et al., 2013. Europe needs a new vision for a Natura 2020 e11842. network. Conservation Letters, 6 (6), 462-467. Consoli, P., Martino, A., Romeo, T., Sinopoli, M., Perzia, P. et Hooper, D.U., Adair, E.C., Cardinale, B.J., Byrnes, J.E., Hun- al., 2015. The effect of shipwrecks on associated fish as- gate, B.A. et al., 2012. A global synthesis reveals biodiver- semblages in the central Mediterranean Sea. Journal of the sity loss as a major driver of ecosystem change. Nature, 486 Marine Biological Association of the United Kingdom, 95 (7401), 105-108. (1), 17-24. IUCN, 2015. https://www.iucn.org/. (Accessed 13 October Consoli, P., Esposito, V., Battaglia, P., Altobelli, C., Perzia, P. et 2020). al., 2016. Fish distribution and habitat complexity on banks Italian Committee of IUCN, 2016. http://www.iucn.it/. (Ac- of the Strait of Sicily (Central Mediterranean Sea) from re- cessed 13 October 2020). motely-operated vehicle (ROV) explorations. PLoS ONE, Koslow, J.A., 1996. Energetic and life‐history patterns of deep‐ 11(12), e0167809. sea benthic, benthopelagic and seamount‐associated fish. Consoli, P., Andaloro, F., Altobelli, C., Battaglia, P., Campa- Journal of Fish Biology, 49, 54-74. gnuolo, S. et al., 2018. Marine litter in an EBSA (Ecologi- Kvile, K.Ø., Taranto, G.H., Pitcher, T.J., Morato, T., 2014. A cally or Biologically Significant Area) of the Central Med- global assessment of seamount ecosystems knowledge us- iterranean Sea: abundance, composition, impact on benthic ing an ecosystem evaluation framework. Biological Con- species and basis for monitoring entanglement. Environ- servation, 173, 108-120. mental Pollution, 236, 405-415. Lodolo, E., Ben-Avraham, Z., 2015. A submerged monolith in CBD-COP 12 Decisions, 2014. Twelfth meeting of the Confer- the Sicilian Channel (central Mediterranean Sea): Evidence ence of the Parties to the Convention on Biological Diver- for Mesolithic human activity. Journal of Archaeological sity (CBD), www.cbd.int/decisions/cop/?m=cop-12 (Ac- Science: Reports, 3, 398-407. cessed 13 October 2020). Lodolo, E., Sanfilippo, R., Rajola, G., Canese, S., Andaloro, Costello, M.J., Coll, M., Danovaro, R., Halpin, P., Ojaveer, H. F. et al., 2017. The red coral deposits of the Graham Bank et al., 2010. A census of marine biodiversity knowledge, area: Constraints on the Holocene volcanic activity of the resources, and future challenges. PLoS ONE, 5 (8), e12110. Sicilian Channel. GeoResJ, 13, 126-133. Council of Europe, 1979. Explanatory Report Concerning the Molinari Novoa, E.A., Guiry, M.D., 2020. Reinstatement of the Convention on the Conservation of European Wildlife and genera Gongolaria Boehmer and Ericaria Stackhouse (Sar- Natural Habitats, Bern Convention: Convention Opened gassaceae, Phaeophyceae). Notulae Algarum, 171, 1-10.

Medit. Mar. Sci., 22/2 2021, 297-316 315

http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 06/10/2021 09:44:35 | Moser, D., Ellmauer, T., Evans, D., Zulka, K.P., Adam, M. et Rosso, A., Di Martino, E., 2016. Bryozoan diversity in the Med- al., 2016. Weak agreement between the species conserva- iterranean Sea: an update. Mediterranean Marine Science, tion status assessments of the European Habitats Directive 17 (2), 567-607. and Red Lists. Biological Conservation, 198, 1-8. Sinopoli, M., Consoli, P., Perzia, P., Romeo, T., Andaloro, F., Orellana, S., Hernández, M., Sansón, M., 2019. Diversity of 2015. Distribution of fish fauna associated with a shipwreck Cystoseira sensu lato (Fucales, Phaeophyceae) in the east- in the southern Tyrrhenian Sea: vertical distribution and ern Atlantic and Mediterranean based on morphological and shipwreck structures. Journal of Applied Ichthyology, 31, DNA evidence, including Carpodesmia gen. emend. and 96-101. Treptacantha gen. emend. European Journal of Phycology, Staudigel, H., Koppers, A.A., Lavelle, J.W., Pitcher, T.J., 54 (3), 1-19. Shank, T.M., 2010. Defining the word “seamount”. Ocean- Pérès, J.M, Picard, J., 1964. Nouveau manuel de bionomie ben- ography 23 (1), 20-21. thique de la Méditerranée. Recueil des travaux de la Station UNCLOS, 1982. UNCLOS, Montego Bay Convention. https:// Marine d’Éndoume, 31 (47), 1-137. www.un.org/depts/los/convention_agreements/texts/un- Pitcher, T.J., Morato, T., Hart, P. J., Clark, M.R., Haggan, N. et clos/unclos_e.pdf (Accessed 13 October 2020). al., (Eds), 2007. Seamounts: Ecology, Fisheries, and Con- UNEP, 1973. The Convention on International Trade in Endan- servation, vol. 12. Blackwell Publishing, Oxford, UK, 527 gered Species of Wild Fauna and Flora (CITES, Washing- pp. ton Convention). https://www.cites.org/ (Accessed 13 Oc- Pons-Fita, A., Verdura, J., Santamaría, J., Kersting, D.K., Bal- tober 2020). lesteros, E., 2020. Coexistence of the reef-building coral UNEP, 1992. The Convention on Biological Diversity (CBD). Cladocora caespitosa and the canopy-forming alga Trep- https://www.cbd.int/convention/ (Accessed 13 October tacantha ballesterosii: Description of a new Mediterranean 2020). habitat. Scientia Marina, 84 (3), 263-271. UNEP/MAP, 1995. Protocol on Specially Protected Areas and RAC/SPA, 2008. Joint Management Action of the European Biological Diversity in the Mediterranean (SPA/BD pro- Community with the United Nations Environment Pro- tocol) of Barcelona Convention 1995. http://www.rac-spa. gramme/Mediterranean Action Plan. Regional Activity org/sites/default/files/protocole_aspdb/protocol_eng.pdf Centre for Specially Protected Areas (RAC/SPA). http:// (Accessed 13 October 2020). www.rac-spa.org/node/597. (Accessed 13 October 2020). UNESCO, 2013. Final report of the 25th Session of the Inter- Relini, G., Giaccone, G. (Eds), 2009. Gli habitat prioritari del national Coordinating Council of the Man and the Bio- protocollo SPA/BIO (Convenzione di Barcellona) presenti sphere (MAB) Programme. Available at: http://www. in Italia: schede descrittive per l’identificazione. Erredi, unesco.org/new/fileadmin/MULTIMEDIA/HQ/SC/pdf/ Grafiche Editoriali, Genova, 367 pp. SC-13-CONF-225-11_Final_Report_en.pdf Rockström, J., Steffen, W., Noone, K., Persson, Å., Chapin III, Worm, B., Barbier, E.B., Beaumont, N., Duffy, J.E., Folke, C. F.S. et al., 2009a. A safe operating space for humanity. Na- et al., 2006. Impacts of biodiversity loss on ocean ecosys- ture, 461 (7263), 472-475. tem services. Science, 314 (5800), 787-790. Rockström, J., Steffen, W.L., Noone, K., Persson, Å., Chapin Würtz, M., Rovere, M., (Eds), 2015. Atlas of the Mediterranean III, F.S. et al., 2009b. Planetary boundaries: exploring the Seamounts and Seamount-like Structures. IUCN, Gland, safe operating space for humanity. Ecology and Society, 14 Switzerland and Málaga, Spain, 276 pp. (2), 1-33.

Supplementary data

The following supplementary information is available online for the article:

Appendix S1: List of taxa detected on the Graham (Gr), Nereo (Ne) and Pantelleria Vecchia (Pa) banks through ROV visual assessment and ROV grabber sample.

316 Medit. Mar. Sci., 22/2 2021, 297-316

http://epublishing.ekt.gr | e-Publisher: EKT | Downloaded at 06/10/2021 09:44:35 |